Revue suisse de Zoologie 1 17 (4): 771-800; decembre 2010
The Psilidae (Diptera, Acalyptrata) of Switzerland,
with description of two new species from Central Europe
Anatole I. SHATALKIN 1 & Bernhard MERZ 2
1 Zoological Museum, Moscow State University, B. Nikitskaja str. 6, K-9,
103009 Moscow, Russia.
E-mail: shatalkin@zmmu.msu.ru
2 Museum d’histoire naturelle Geneve, C. P. 6434, 121 1 Geneve, Switzerland.
E-mail: bernhard.merz@ville-ge.ch
The Psilidae (Diptera, Acalyptrata) of Switzerland, with description of
two new species from Central Europe. - The family Psilidae is reviewed
for Switzerland and data are listed for all species currently known from this
country. Chamaepsila sardoa (Rondani) and Ch. unilineata (Zetterstedt) are
recorded for the first time from Switzerland. Psila (s. str.) Helvetica sp. n.
(Switzerland) and Chamaepsila conjusa sp. n. (France, Switzerland) are
described and illustrated. Keys for the European species of Psila s. str.
Meigen and the Chamaepsila pallida group are provided. A summary of
diagnostic characters of the genera of Psila s. lat. is given. An updated
checklist of the 36 Swiss species is presented.
Keywords: Psilidae - Switzerland - new species - keys - checklist.
INTRODUCTION
The Psilidae are a small family of acalyptrate Diptera with 335 species
described from all biogeographical regions. Most species occur in the Holarctic Region
with 153 species currently known from the Palaearctic Region. They are considered to
be monophyletic with some well-supported apomorphic character states: wing
venation peculiar (the costa has a break at the distal end of the subcosta which is
reaching the costa as a fold; the distal margin of cell cup (anal cell, posterior cubital
cell) is closed by a straight vein CuA2) and chaetotaxy reduced (0-1 notopleural setae;
acrostichal setulae usually not differentiated; pleuron bare or covered with setulae, but
without setae).
Traditionally, the family is divided into two subfamilies, the Chylizinae and the
Psilinae. The Chylizinae (116 species worldwide, particularly numerous in the
Afrotropical and Oriental Regions, 22 species in the Palaearctic Region) are probably
plesiomorphic whereas the Psilinae (216 species described mainly from the Holarctic
Region, few species in the Afrotropical and Oriental Regions, unknown in the
Neotropics) are usually considered to be a monophyletic group. The Psilinae consist
mainly of slender species ranging from 3 mm (often in Psila s. lat.) to 13 mm (some
Loxocera spp.). They have usually a triangular head with a receding face in profile.
Manuscript accepted 09.06.20 10
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A. I. SHATALKIN & B. MERZ
Their parafacialia lacks silvery white microtrichia. The male terminalia are characteri-
zed by well-developed parameres and by the absence of a surstylus. The Chylizinae are
more stumpy flies and differ from the Psilinae by the developed postcoxal bridge and
the presence of a surstylus in the male. Their parafacialia is silvery microtrichose.
McAlpine (1997) considered this last character as groundplan condition of Diopsoidea
(Diopsidae and Syringogastridae), but it is also present in some species of other fami-
lies (Megamerinidae, Strongylophthalmyiidae, Tanypezidae).
Recently, Shatalkin (2001) proposed a third subfamily, the Belobackenbardiinae
for three species of Belobackenbardia Shatalkin from South Africa. They are externally
similar to primitive species of Psila s. lat., but their parafacialia is silvery micro-
trichose. Also, their male terminalia do not have any resemblance with those of other
Psilidae and related families. It is possible that their similarity with Psila s. lat. is due
to convergence.
The known larvae of Psilidae are phytophagous (Ferrar, 1987; Iwasa, 1998).
They are tunneling in stems, rootstocks and bulbs, or live under the bark of thin twigs
of trees. The larvae of the common European species Chyliza leptogaster (Panzer) may
cause phloem necroses in deciduous trees (Dengler, 1997). Those of Chyliza annulipes
Macquart have been recorded from conifers, in particular from resinous wounds
(Lyneborg, 1987). Chyliza vittata has been reared from mines in stems and roots of
Orchidaceae (de Meijere, 1940; Petit, 1982) whereas the larvae of the externally
similar Chinese Chyliza bambusae Yang & Wang develop in roots of bamboo (Yang &
Wang, 1988). The Nearctic Chyliza leguminicola Melander is feeding from the lower
part of Lupinus polyphyllus (Melander, 1920). Species of Loxocera and some Psila s.
lat. are often found in marshland where their larvae are developing in stems of
Cyperaceae ( Carex ) or Juncaceae (Juncus).
Only few species of the family are of economic importance. The larvae of the
carrot fly, Chamaepsila rosae (Fabricius), may cause important damage in edible roots
of carrots and other umbelliferous crops (celery [= Apium graveolens\ , parsley
[= Petroselinum hortense], parsnip [= Pastinaca sativa]) and some cruciferous species
(cabbage [= Brassica sp.], turnip [= Brassica rapa]). Another pest species is
Chamaepsila nigricornis (Meigen) whose larvae feed in roots of ornamental plants.
REVIEW OF THE FAUNISTIC RESEARCH IN SWITZERLAND
Switzerland is a small country in Central Europe but of a remarkable diversity
with currently almost 7'000 species of Diptera recorded (Merz et al. , 2007). This may
be explained by the presence of species which are widely distributed in temperate
Central Europe living in various types of grassland from swamps to rather dry mea-
dows and in deciduous and evergreen forests. In addition, a notable influence of the
Mediterranean fauna may be recorded in Southern Switzerland and the region of
Geneva. The dry Central Alpine valleys are famous for some steppose. East European
species. Finally, numerous Scandinavian or boreal species were found in Subalpine and
Alpine Regions of the Alps and the Jura chain.
The Psilidae of Switzerland have never been studied. The only historical list
with six reliable records was published by Ringdahl (1957). Later, Merz (1998)
established the Swiss checklist with 31 species based on about 600 specimens housed
PS I LI DAE OF SWITZERLAND
773
in the 10 most important public collections of Switzerland. However, no data with
precise information (localities, dates of collecting) were provided. One species,
Loxocera maculata , was later added by Merz et al. (2002).
The motivation of the present paper is threefold. First, the knowledge of the
Swiss fauna is reviewed with inclusion of distributional data for all species and a
revised checklist is provided here in the appendix. Second, the occasion is taken to
describe two species new to science which were discovered during the preparation of
the paper. And finally, updated keys are given for some taxonomically difficult groups
in the light of new results which were obtained by the study of the present specimens.
MATERIAL AND METHODS
This study is based on about 330 specimens which were almost exclusively
collected in Switzerland (CH) and which are deposited in the collection of the Natural
History Museum Geneva (MHNG). Few duplicate specimens are also kept in the
Zoological Museum of the Moscow State University (ZMUM). In addition, the type
series of one species, Scatophaga pallida Fallen, deposited in the Swedish Museum of
Natural History (NRS) was examined.
The responsabilities are distributed as follows: The first author identified at
least one specimen of each species. He prepared a draft version of the manuscript with
the descriptions and illustrations of the species and with the elaboration of the new
facts and keys arising from the study of the specimens. The second author identified
further specimens of almost all species and he prepared an updated version of the
manuscript.
For identification, the following keys and revisions were used: Hennig (1941),
Collin (1944), Lyneborg (1964), Shatalkin (1986), Wang (1988). Greve & Midtgaard
(1989), Shtakel'berg (1989), Carles-Tolra (1993a), van der Goot & van Veen (1996),
Iwasa (1998), and Greve & Skartveit (2000). Data about the distribution of the species
in Europe is partly taken from Pape (2007).
Terminology and nomenclature in the systematic part follows basically Iwasa
(1998) and Merz & Haenni (2000). The genera are arranged in a phylogenetic order
and the species alphabetically within the genera. The specimens are listed in alphabe-
tical order for cantons.
SYSTEMATIC PART
Subfamily Chylizinae
This subfamily includes the single genus Chyliza. It is characterized by the
following unique combination of characters: anatergal area of laterotergite enlarged,
callus-like; occiput concave; face nearly perpendicular, not retreating; head not trian-
gular in profile; usually three (pairs of) scutellar setae; postcoxal bridge developed;
anal cell (posterior cubital cell, cup) shorter than 2nd basal cell.
Genus Chyliza Fallen, 1820
Type species: Musca leptogaster Panzer, 1798, by subsequent designation of
Westwood, 1840: 146.
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A. I. SHATALKIN & B. MERZ
Comments: In Europe, five species are recognized (Hennig, 1941; Soos, 1984)
which all occur in Switzerland. "Chyliza" gracilis Loew, 1854, is considered here to
belong to the subfamily Psilinae (see below).
Chyliza annulipes Macquart, 1835
Material Switzerland: Id, GE, Corsier-Port, vitre veranda, 27-31 V 2005, leg. C.
Besuchet. -Id. ZH, 400 m, Embrach-Haumiili. 9 V 1998, leg. B. Merz. - 1 d , 1 9 , ZH, 460 m,
Zurich-Ziegelhiitte, 10 V 1997, leg. B. Merz.
Chyliza extenuata (Rossi. 1790)
Material Switzerland: 1 d , 1 9 . FR, 460-650 m. Mt. Vully, Bas Vully, 6 VI 2003, leg.
B. Merz & F. Amiet. - Id, GE, 510 m, Bemex. Signal, 24 V 2002, leg. B. Merz. - Id, GE,
Corsier-Port, vitre veranda, 1-31 V 2009, C. Besuchet.
Comments: Larvae develop in roots of broomrapes ( Orobanche spp.). Outside
of Europe this species is recorded from Central Asia.
Chyliza leptogaster (Panzer, 1798)
Material Switzerland: Id, GE, Corsier-Port, vitre veranda, 20-30 VI 2008, leg. C.
Besuchet. - 29 , GE, 350 m, Chancy, La Laire, 1 VII 2001, leg. B. Merz. -19, GE. 500 m,
Bernex-Signal, 17 VI 2009, B. Merz. -Id, GR, 980 m, Andeer-Clugin, 17 V 1994, leg. B. Merz.
-19, SG. 610 m, Wattwil. 11 VII 1997, leg. B. Merz. - Id, TI, 350 m, Biasco-Loderio,
7180/1375, 16 V 2006, leg. B. Merz. - 19, TI, 600-1100 m, Monte San Giorgio,
717180/085700. 5 VII 2001 , leg. B. Merz. -19. VS, 750 m. Branson/Follateres, 9 VI 2004, leg.
B. Merz & J.-P. Haenni. -19, VS, 700-770 m, Raron-Heidnischbiel, 26 V 1999, leg. B. Merz.
- 1 9 , VS, 750 m, Raron-Heidnischbiel, 3 VII 2003, leg. Merz, Smit & van Det. - 1 d , VS, 625
m, Leuk-Platten, 8 VI 2001 . leg. B. Merz & B. Landry. -19. ZH, 650 m, Zurich-Ziirichberg, 26
VII 1995, M.Fal [Malaise trap], S.Ungricht.
Chyliza nova Collin, 1944
Material Switzerland: 1 9 , GE, 440 m, Bernex-Saule, 14 VI 2009, B. Merz. -19,
GE, 440 m, Bernex-Saule, 19 VI 2008, leg. B. Merz. -19, GE, 440 m, Bernex-Signal, 25 VI
2009. B. Merz. -Id, GE, Corsier-Port, vitre veranda, 1-30 VI 2004, leg. C. Besuchet. -19,
VD, Bonvillars, La Coudre, 10 VII 1991 , leg. J. Steffen. -Id, ZH. 450 m, Zurich-Waldgarten,
24 V 1996. leg. B. Merz. - Id. ZH. 460 m, Zurich-Ziegelhiitte, 29 VI 1997, leg. B. Merz.
Chyliza vittata Meigen, 1826
Material Switzerland: 1 9 . BE, 650-950 m, Mt. Raimeux. Corcelles. 2 VI 2003, leg.
B. Merz. -Id, GE, 350 m, Cartigny, Moulin de Vert, 24 V 1999, leg. Merz & Muller.
Subfamily Psilinae
Tribe Loxocerini
Comments: Traditionally, this tribe includes Psila-Yike flies which are charac -
terized by a long postpedicel which is longer than the head, and by a pad of short
setulae which are often present near the ventral tip of the hind femur. In Europe, this
tribe is represented by the genus Loxocera s. lat. (see below). Shatalkin (1998) pro -
posed a new subgenus, Asiopsila Shatalkin. for Oriental species of small size (3.5-5
mm) without the pad of short setulae on the hind femur but with a whitish spot in the
upper part of the katepistemum. Later, Buck & Marshall (2006b) transferred this sub-
genus in the tribe Psilini where it is considered to be a subgenus of Psila s. str. Their
PSIUDAE OF SWITZERLAND
775
conclusion seems to be justified. However, it should be noted that the structure of the
male terminalia with the reduced parameres in Asiopsila differs strongly from the
superficially similar external morphology in other taxa of the Psilini. In this respect
they resemble species of Loxocera s. lat. Following this conclusion Loxocera glandi-
cula Iwasa, 1993, which is close to L. aristata , is the only Oriental species of Loxocera
s. str.
Hennig (1941) and Steyskal (1987) treated Platystyla Macquart, 1835, as a sub-
genus of Loxocera . Species of Platystyla are characterized by extremely long antennal
segments and a strongly thickened and flattened arista which is inserted near the
middle of the postpedicel or even more apically (the part distal of the insertion of the
arista on the postpedicel is 1 .0-1.3 times as long as the part basal of it). Earlier, Frey
(1925) treated Loxocera and Platystyla as distinct genera and he proposed the new sub-
genus Imantimyia Frey within Loxocera for Loxocera albiseta (Schrank) and
morphologically similar species (L. sylvatica and L. jidviventris). Recently, Buck &
Marshall (2006b) proposed a new classification based on a phylogenetic study of
Nearctic and Palaearctic Loxocerini, in particular of the egg structure and the male
terminalia. They concluded that Loxocera sensu Frey (1925) or Hennig (1941) is para-
phyletic and they suggested that Imantimyia is the sister-group of {Loxocera s. str. +
Platystyla). As a consequence of this phylogenetic study they treat Imantimyia and
Platystyla as subgenera of Loxocera. However, based on its morphological peculia-
rities with the strongly modified antennae it is difficult to accept the proposition that
Platystyla with its 5 Oriental and Palaearctic species (Frey, 1955) should be considered
as subgenus of Loxocera. In order to preserve the new phylogenetic hypothesis of Buck
& Marshall (2006b) the following concept is proposed here: Imantimyia is treated as a
separate genus, whereas Platystyla is placed as subgenus within Loxocera. Following
this proposition it should be noted that Tropeopsila Shatalkin, 1983, described for two
East Palaearctic species and which was not studied by Buck & Marshall (2006b), is a
subgenus of Loxocera.
Species of Loxocera and Imantimyia develop in stalks of sedges (de Meijere,
1941; Chandler, 1975; Ferrar, 1987). The tribe contains currently 54 species; 18 species
are known from the Palaearctic Region. All 7 European species (Pape, 2007) occur in
Switzerland.
Genus Loxocera Meigen, 1803
Type species: Musca aristata Panzer, 1801, by monotypy.
Comments: In the current interpretation, Loxocera is a well characterized
genus. It differs from the other genus of the tribe, Imantimyia , by the following
characters (see also Buck & Marshall, 2006b): frons with a pair of desclerotized
velvety vittae; lunule semicircular between antennal bases and anterior margin of
frons, crest-like (usually hidden in Imantimyia); alula bare at least medially; male
pregenital sclerite large and setulose (small and bare in Imantimyia); female cerci not
fused with tergite 10 (fused in Imantimyia).
According to the present concept, the genus contains besides the nominal sub-
genus also the two subgenera Platystyla Macquart and Tropeopsila Shatalkin. In
Europe, two species of Loxocera s. str. and one species of Platystyla are known, all
occurring also in Switzerland.
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A. I. SHATALKIN & B. MERZ
Subgenus Loxocera Meigen, 1803
Loxocera (s. str.) aristata (Panzer, 1801)
Material Switzerland: 1 9 , BE, 995 m, Tramelan, La Tourbiere, 4 VI 2003, Merz,
Haenni & Rapp. -19, GE, 400 m, Bemex, Chante-Merle, 1 VI 2009, leg. B. Merz. -19, GE,
420 m, Bemex, Bois des Mouilles, 19 VIII 2006, leg. B. Merz. -19, GE, Corsier-Port, vitre
veranda. 1-15 IX 2008, C. Besuchet. - 1 6 , GE, Corsier-Port, vitre veranda, 27-31 V 2005, leg.
C. Besuchet. -19, GL, 1500 m, Pragelpass, 5 VIII 1991, leg. Bachli, Beuk, Merz. -19, JU,
1020 m, Les Breuleux, La Tourbiere, 4 VI 2003, leg. Merz, Haenni & Rapp. -19, JU, 1000 m,
La Chaux-des-Breuleux, La Tourbiere, 28 VI 2003, leg. B. Merz. - 1 9, VS, 1700-1900 m,
Morgins, La Chaux-Culet, 21 VI 2003, leg. B. Merz. -19, VS, 1700-1900 m, Morgins-En Tey,
22 VI 2003, leg. B. Merz.
Loxocera (s. str.) maculata Rondani, 1876 Figs P1-P2
Material Switzerland: 4d , 5 9 , VS, 2000 m, Simplon/Hospiz, 645450/121680, 4 VII
2001 , leg. B. Merz.
Comments: The status of this species is subject to different interpretations.
Some authors synonymized L. maculata with L. aristata , for instance Hennig (1941)
and Soos (1982) who studied the types in the collection of Rondani. Later, however,
Soos (1984) treated L. maculata as a good species with L. atriceps Bigot, 1886, and
L. yerburyi Austen, 1899, as synonyms. This concept was adopted by Schacht (1996)
for Bavaria, van der Goot & van Veen (1996) for the Netherlands and Merz et al.
(2002) for Switzerland. On the other hand, Bartak & Carles-Tolra (2006) in the
Catalogue of Diptera of Czech and Slovak Republics listed L. maculata as subspecies
of L. aristata.
L. atriceps , considered to be a synonym of L. maculata by Soos (1984), is now
believed to be a variety of L. aristata differing only in its colouration.
The Swiss specimens of L. maculata differ from those of other countries. They
are characterized by a rather short postpedicel which is shorter than the length of the
frons (measured from the posterior ocelli to the lunule), by a long arista which is longer
than the postpedicel, and by an entirely black thorax (Figs PI , P2).
In external characters L. maculata differs from L. aristata by its almost black
thorax including the scutellum. On the other hand, no clear differences in the structure
of the male terminalia have yet been found. Therefore, additional specimens from
various localities are necessary in order to clarify the taxonomy of the two taxa. For
the time being they are treated as distinct species.
Subgenus Platystyla Macquart, 1835
Type species: Loxocera hoffmannseggi Meigen, 1826, by monotypy.
Comments: In the current interpretation the subgenus contains five species, four
of which are recorded from the East Palaearctic Region (Russian Far East, Japan,
China). One species occurs in Europe.
Loxocera (Platystyla) hoffmannseggi Meigen, 1826
Material Switzerland: 1 6 . Bern, Kirchenfeld, 19 VIII 1924, leg. Th. Steck. -Id, ZH,
410 m, Embrach-Haumlili, 20 VIII 1997, leg. B. Merz.
Genus Imantimyia Frey, 1925
Type species: Nemotelus albiseta Schrank, 1803, by original designation.
PSILIDAE OF SWITZERLAND
111
Plate 1-6
Habitus and details of Psilidae. (PI, P2) Loxocera maculata Rondani, habitus lateral and dorsal
(CH, VS, 2000 m, Simplon/Hospiz, 645450/121680, 4 VII 2001). (P3, P4) Psila Helvetica sp. n.
(holotype, CH, VS, 650 m, Leuk-Pfynwald, 23 IX 1992). (P5) Chamaepsila pallida (Fallen)
(male syntype, probably from Sweden, Esperod, NRS). (P6) head of Ch. pallida (female
syntype, probably from Sweden, Esperod, NRS).
Comments: The European fauna contains four species which are all known
from Switzerland.
Imantimyia albiseta (Schrank, 1803)
Material Switzerland: 4 9 , GE, 420 m, Bernex, Bois des Mouilles, 19 VIII 2006, leg.
B. Merz. - 1 9 , SG, 910 m, Unterwasser, 15 VIII 1997, leg. B. Merz. - 1 6 , ZH, 450 m, Zurich-
Allmend, 10 IX 1996, leg. B. Merz.
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A. I. SHATALKIN & B. MFJRZ
Imantimyia fulviventris (Meigen, 1826)
Material Switzerland: 1 8 , 1 9 , ZH, 515 m, Illnau-Wildert, 5 VIII 1997, leg.
B. Merz.
Comments: Both specimens were collected at light in a peat-bog.
Imantimyia nigrifrons (Macquart, 1835)
Material Switzerland: 1 8 , GE, 360 m, Dardagny, Le Moulin, 8 VII 2002, leg. B.
Merz.
Imantimyia sylvatica (Meigen, 1826)
Material Switzerland: \ 8 , VS, 1700-1900 m, Morgins, La Chaux-Culet, 21 VI
2003, leg. B. Merz. -18, ZH, 410 m, Glattfelden-Bahnhof, 1 5 V 1993, leg. B. Merz. - 1 9 , ZH,
620-670 m, Zurich-Ziirichberg, 11 V 1995, leg. B. Merz.
Tribe Psilini
Comments: This is the most diverse tribe of the family in Europe with 34
described species. It includes notably Psila Meigen, 1803, with its confusing history.
It was described for species which could not be placed elsewhere and which exhibit a
series of negative characters. For instance, the type-species, Musca fimetaria Linnaeus,
is characterized by numerous reductions in its chaetotaxy. With increasing knowledge
of the group, also from other biogeographical areas than the Western Palaearctic
Region, numerous generic or suprageneric taxa were proposed which were later often
synonymized or given only subgeneric status within Psila Meigen. Such a phenetic
approach was justified before the development of the principles of phylogeny. In all
cases the status of several species remained unresolved and resulted in residual taxa.
Hendel (1917) described the genus Chamaepsila to accommodate species of Psila with
postocellar setae developed, and thus differing from Psila s. str. where they are absent
(e. g., the type-species M . fimetaria) . Few years later, Frey (1925) proposed the new
genus Oxypsila for Psila abdominalis Schummel which lacks dorsocentral and post-
ocellar setae. The remaining species of Psila s. str. are thus a morphologically distinct
group which may be regarded as a specialized offshoot of the more generalized genus
Chamaepsila. However, it will be shown below that even the latter genus should be
considered as a residual taxon including species of ambiguous affinities.
Hendel (1934) described the genus Synaphopsila for one Chinese species, S.
hummeli Hendel, 1934, which is characterized by the presence of three vertical setae
instead of two setae present in Oxypsila. Later, Wang & Yang (1996) synonymized the
two genera and included S. hummeli and three new Chinese species in Oxypsila. While
studying S. hummeli , new differences compared with P. abdominalis were found, like
the absence of notopleural setae (as it is also the case in Psilosoma Zetterstedt, but one
notopleural seta is present in P. abdominalis'). There are evidences that S. hummeli (and
further 9 East Palaearctic species of the same group) does not form a monophylum with
P. abdominalis. Synaphopsila is probably closer related to Psila s. str. and therefore
considered here to be a subgenus of the latter.
Frey (1925) proposed the genus Tetrapsila for Psila obscuritarsis Loew, 1856,
a species with four scutellar setae (two pairs), which was formerly placed in
PSIUDAE OF SWITZERLAND
779
Chcimciepsila whose species have only one (pair of) apical scutellar setae. Later, Seguy
(1936) added Tetrapsila longipennis Seguy, 1936, from the Azores Islands to the
genus. This species, however, belongs in our opinion to Afropsila Shatalkin, and its
inclusion in Tetrapsila was based on morphological similarity. It is characterized by
black halteres in the female and some other features which are typical for Afropsila.
The number of scutellar setae is not of generic value, as two pairs are also present in
" Chyliza " gracilis Loew (see below) and the Asian Xenopsila Buck (Buck & Marshall,
2006a).
Zetterstedt (1860) described the genus Psilosoma for the two species
Scatophaga audouini Zetterstedt, 1835, and Scatophaga lefebvrei Zetterstedt, 1835.
The status of this genus, however, is not established. Some authors, like Hennig (1941),
Soos (1984) or Pape (2007), considered it as a good genus different from Psila s. lat.
But Collin (1944) expressed some doubts "whether these two species are generically
distinct" from Psila and he treated Psilosoma as subgenus of Psila. This opinion was
followed by Shatalkin (1986, 1989) and Iwasa (1998). Psilosoma differs from Psila
and Chamaepsila by the absence of notopleural and supra-alar setae, the expanded and
curved hind femur in the male, and the laterally compressed ovipositor in the female.
The male terminalia are also peculiar with their long, dilated and poorly sclerotized
aedeagus (Shatalkin, 1986. plate 1, figs 3-4). Notopleural setae are also absent in
Synaphopsila Hendel. The two genera differ in addition to some external characters
also in their distribution. Psilosoma is an European genus whereas species of
Synaphopsila occur in the Russian Far East, China, Korea and Japan. Pending a
cladistic analysis of this group of genera we are following the tradition of European
dipterists and treat Psilosoma in the present paper as a good genus.
Hennig (1941) described Psila problematica from Germany. However, a com-
parison of its type with that of Chamaepsila sibirica Frey, 1925, showed that they are
conspecific and they were synonymized by Shatalkin (1986). He proposed for this
species a new subgenus in Psila , Freyopsila Shatalkin, 1986, in order to accommodate
species of Psila without postocellar setae. This group contains currently eight species
which are mainly distributed in the Eastern part of the Palaearctic Region.
Another problem concerns American species of Psila s. lat. with an elongated
postpedicel for which Johnson (1920) described the genus Pseudopsila (type species:
Loxocera fallax Loew, 1869). Later, Frey (1955) described two Oriental species from
Myanmar ( Pseudopsila maculipennis and P. nigricollis) and Shatalkin (1983) added
the first Palaearctic species to the genus (P. nigrifidva). A careful study of the types of
the two species described by Frey and the Nearctic Pseudopsila collaris (Loew, 1869)
and P. lateralis (Loew, 1860) revealed that P. nigrifidva does not belong to this genus
but seems to be closer related to Psila sibirica (Frey), and it was transferred to
Freyopsila (Shatalkin. 1983). More recently, Buck & Marshall (2006a) revised the
Nearctic species of Pseudopsila and concluded based on a cladistic analysis that the
type species of Pseudopsila (L. fallax), as well as two other species, should be syno-
nymized with Psila s. str. and proposed Xenopsila as new subgenus of Psila for the
other Nearctic species previously placed in Pseudopsila (P. bivittata , P. collaris , P.
lateralis) as well as for the Oriental and Palaearctic species (P. (X.) arbustorum
Shatalkin, P. (X.) nemoralis Shatalkin, P. ( X .) tetrachaeta Shatalkin).
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A. I. SHATALKIN & B. MERZ
Among European Psila s. lat. a short comment should be made for Chyliza gra-
cilis Loew, 1854, which is characterized by a long, finely setulose, white arista.
Freidberg & Shatalkin (2009) presented a review of its confusing taxonomic history.
After a careful study of six specimens from Cyprus, Israel and Turkey they concluded
that this species should be transferred in Chamaepsila thus becoming a junior homo-
nym of Psila gracilis Meigen. 1826 which is now considered to be a synonym of
Chamaepsila buccata (Fallen, 1826). Therefore, they proposed the replacement name
Chamaepsila setalba Freidberg & Shatalkin, 2009 for Chyliza gracilis Loew, 1854.
However, its combination of characters differs from other species of Chamaepsila and
it represents probably a separate lineage within the genus. Moreover, a single female
from Bulgaria (Sandanski) differs in some points from Ch. setalba and may represent
another yet undescribed species. Therefore, further studies based on more specimens
are needed in order to propose well-funded arguments for the generic position and for
the intra-specific variability of the included taxa.
A long-lasting debate is dividing students of the family Psilidae. Whereas some
authors consider the various taxa of Psila s. lat. as valid genera (Hennig, 1941; Soos,
1984; Pape, 2007), others treat them as subgenera of Psila s. lat. (Shatalkin, 1989,
2008; Iwasa, 1998). Based on a morphological analysis of external characters alone
this question cannot be resolved. The differences ("gaps") between the taxa are
weightened in a different manner by each student, and apparently "good" characters in
one biogeographical region may be unreliable in other areas because of intermediate
species or species-groups. For instance, the Non-European taxon Xenopsila described
as subgenus of Psila (Buck & Marshall, 2006a) is characterized by a most peculiar type
of male terminalia with a long, thin, scerotized aedeagus. But some species of
Loxocera , in particular the L. ( Imantimyia ) achaeta group (including the Nearctic
species L. cylindrica Say, L.fumipennis Coquillett, L. ignyodactyla Buck, the Japanese
L. lutulenta Iwasa and the Palaearctic L. achaeta Shatalkin and L. nigrifrons Macquart)
are similar and overlap with those of Xenopsila.
Species of Freyopsila , Psilosoma , Synaphopsila and ( Chamaepsila + Psila)
form a group which is characterized by a similar, more simple structure of male termi-
nalia. Male terminalia of Psila s. str. and Chamaepsila resemble each other and are
clearly separated from the other genera and represent a third level of the structure of
the male terminalia. If only European species of the two taxa are considered then they
form clearly two distinct groups which would deserve generic rank. However, with the
inclusion of Asian species differences between them become unreliable because of
species with an intermediate combination of characters. For this reason Iwasa (1998)
treated Chamaepsila as a subgenus of Psila s. lat. It is therefore not surprising that the
Japanese species Psila magna (Shatalkin, 1983) was described as Chamaepsila
because it has a well developed postocellar seta as it is the case for other Chamaepsila.
On the other hand, Psila nigripalpis Shatalkin, 1983, is similar to some species of
Chamaepsila , in particular of the Ch. rnfa group, but lacks the postocellar setae and
was described originally in Psila.
Based on these problems all available species of the two taxa were studied again
in detail in order to redefine their generic placements. It was found that a new
character, the presence of fine, inconspicuous setulae on the posterior half of the
PS 1 LI DAE OF SWITZERLAND
781
anepimeron, is a constant character for species of Psila s. str. Such setulae are absent
in all species of Chamaepsila (except for one species, Cliamaepsila fenestralis
Shatalkin, 1983, from Russian Far East) and other taxa of Psila s. lat. For this reason
we propose here to treat Chamaepsila and Psilosoma as distinct genera and exclude
them from Psila s. str. Some characters which were found to be of diagnostic value for
the various taxa of Psila s. lat. are summarized in Tab. 1 .
Tab. 1. Diagnostic characters of Psila s. lat. In ( ): exceptions, see species listed below
Characters
Taxa
Notopleural
setae
Anepimeron
with [s]etulae
or [b]are
Postocellar
setae
Dorsocentral
setae
Vertical
setae
Scutellar
setae
[pairs]
colour
Arista
y=yellow
w= white
Postpedicel
[l] ong as in
Loxocera
[m] edium,[s]hort
Psila s. str. spp.
1
(b), s
0, 1
(0), 1
2-3
1
(w),y
(m), s
P. abdominalis
1
s
0
0
2
1
y
s
P. crassula , P. magna
1
s
1
1
2
1
y
s
P.fallax
1
b
1
1
2
1
w
m
Chamaepsila spp.
1
b,(s)
1
1-4
2-3
1-2
(w),y
(m), s
Ch. fenestralis
1
s
1
1
2
1
y
s
Ch. obscnritarsis
1
b
1
1
3
2
y
s
Ch. setalba
l
b
1
1
2
1-2
w
m
Afropsila spp.
1
b
0
1
2
1-2
w
s
Asiopsila spp.
1
b
0-1
0-1
2
2
w
1
Frey op si la spp.
1
b
0,(1)
1
2
1
y
1, (m), s
F. maculipennis group
1
b
1
1
2
1
w
m
Psilosoma spp.
0
b
0
0-1
2
1
y
s
Synaphopsila spp.
0
b
0
0-1
2-3
1
w
s
Xenopsila spp.
1
b
1
1
2
2
w
m, s
Genus Psila Meigen, 1803
Type species: Musca fimetaria Linnaeus, 1761, by subsequent designation of
Westwood, 1840: 146.
Diagnosis: Head large, yellow. Postocellar setae absent (present in some
Chinese and Japanese species). Two vertical setae (three in some Chinese species).
Mesonotum with one dorsocentral seta (absent in P. abdominalis). Scutellum with one
(pair of) seta and with soft, scattered, marginal setulae. Anepimeron with soft, fine
setulae in posterior half (also present in Chamaepsila fenestralis Shatalkin, 1983: this
species and Ch. bimaculata Shatalkin, 1983, form an isolated group related to Psila
s. str.).
Distribution: Psila s. str. may be a comparatively young genus. It has a
disjunct distribution in the Palaearctic Region with five West Palaearctic species, one
species from the Kuril Islands and Japan (P. magna Shatalkin), and some species from
the mountains of Western China.
Key to European species of Psila s. str.
la Anepimeron with soft, fine setulae in posterior half; postocellar setae
absent (except for some Asian species); tarsus of mid leg ventrally with
short, black setulae scattered among the golden setulae Psila s. str. 2
782 A. I. SHATALKIN & B. MERZ
lb Anepimeron bare; one postocellar seta present (except for most
specimens of Ch. setcilba Freidberg & Shatalkin and the East Palaearctic
species Ch. nigripalpis ); tarsus of mid leg ventrally with short yellow
setulae Chamaepsilci spp.
2a Postpedicel about 2.5 times as long as its width at base, yellow but dark
infuscated along dorsal margin; abdomen brownish to black; thorax
yellow or reddish yellow with a darker stripe laterally the level of the
supra-alar seta and the dorsal half of the anepisternum; dorsocentral
setae absent. Body length: 6- 1-7 .3 mm. - Europe and Caucasus
Psila ( Oxypsilo ) abdominalis Schummel, 1844
2b Postpedicel at most 1 .5 times as long as its width at base, if yellow then
without darkened dorsal margin; colouration of thorax different; one
dorsocentral seta 3
3a Postpedicel mostly black 4
3b Postpedicel yellow, at most at base of arista with darkened spot 5
4a Mesonotum and subscutellum yellow. Body length: 6.7-7 .2 mm. -
Caucasus Psila (s. str.) caucasica Mik, 1887
4b Mesonotum yellow, but with an unpaired black stripe medially extended
almost to scutellum, and with a pair of black stripes from posterior end
of postpronotal lobe to level of dorsocentral seta (sometimes these black
stripes may be confluent). Subscutellum black. Body length: 6.5-
7.0 mm. - Switzerland Psila (s. str.) Helvetica sp. n.
5a Eye kidney-shaped, distinctly higher than wide; postpedicel at base of
arista with black spot; rays of arista about twice as long as its basal
diameter. Body length: 7. 7-8. 8 mm. - Europe
Psila (s. str.) funetaria (Linnaeus, 1761)
5b Eye roundish, only slightly higher than wide; postpedicel entirely
yellow; rays of arista at most only little longer than its basal diameter.
Body length: 8. 3-8. 5 mm - Europe Psila (s. str.) merdaria Collin, 1944
Subgenus Oxypsila Frey, 1925
Type species: Psila abdominalis Schummel, 1844, by monotypy.
Psila (Oxypsila) abdominalis Schummel, 1844
Material Switzerland: 1 8 , VS, 640 m, Raron, 14 VII 1966, leg. L. & F. Reiser. - 1 $ ,
VS. 623 m, Leuk-Platten, 3 VIII 1998, leg. Merz & Bachli.
Comments: Wang & Yang (1996) described three species from China without
postocellar and dorsocentral setae as Oxypsila ( O . altusfronsa , O. nigricorpa , O. uni-
stripeda ), but without indication of the presence of the soft, fine setulae on the
anepimeron which are characteristic for P. abdominalis and allow its placement in
Psila s. str. (see diagnosis above). Because they were not available for examination
their generic placement remains tentative.
P. abdominalis may be found in mountains of Central and Southern Europe. It
is unknown from the British Isles (Collin, 1944; Chandler, 1998a), Holland (van der
Goot & van Veen, 1996) and Scandinavia. Its northern border passes across Germany,
PSIUDAE OF SWITZERLAND
783
Czech and Slovak Republics, Hungary, and the Caucasus in the East. One specimen,
however, was recently caught in the Moscow Region. It may therefore have a larger
distribution than the few specimens of this rather rarely collected species in the
museum collections indicate.
Subgenus Psila Meigen, 1803
Psila (s. str.) flmetaria (Linnaeus, 1761)
Material Switzerland: 2 9 , BE. 1300 m, Zweisimmen, Gschwand, 7 VI 2003, leg. B.
Merz & M. Eggenberger. -19, GE, 390-430 m, Dardagny, riviere de Roulave, 11 VIII 2000,
leg. Merz & Bachli. -Id, GE. Avusy, Moulin de la Grave, 25 V 2006, leg. H. Boillat. -19,
GR, 1325 m, Ausserferrera, 30 VIII 2006, leg. B. Merz. -19, GR. 1300-1670 m, Ausserferrera-
Cresta, 1 IX 2006, leg. B. Merz. - 29 , JU, 1000 m, La-Chaux-des-Breuleux, La Tourbiere, 28
VI 2003, leg. B. Merz. - 1 9 , SG, 910 m, Unterwasser, 11 VII 1997, B. Merz. - 2 9 , VS, 1900-
2200 m, Visperterminen, Giw-Gebidempass, 21 VII 2004, leg. B. Merz. - 1 9, VS, 1400 m,
Visperterminen, Kreuz, 3 VI 2003, leg. B. Merz. - 1 9 , VS, 1300 m, Morgins-En Tey, 22 VI
2003, leg. B. Merz. -19, ZH, 400 m, Embrach-Haumiili. 24 VI 1995. leg. R. Wunderlin.
Comments: This is one of the most common species of the family in
Switzerland and may be found in various habitats from the lowlands to about 2200 m
in the Alps.
Psila (s. str.) Helvetica sp. n. Figs P3-P4
Holotype: 6 , Switzerland, VS, 650 m, Leuk-Pfynwald, 23 IX 1992, leg. Merz & Otto
(MHNG).
Paratypes: 1 9 , same data as holotype (MHNG). -19, Switzerland, GR, 670-1100 m.
Flims/Bergsturz, 13 IX 1993, leg. B. Merz (MHNG).
Diagnosis: P. Helvetica differs from other European species of Psila s. str. by
the three black stripes on the mesonotum, the black subscutellum, the mostly black
postpedicel. and the short, fine rays on the arista. This combination of characters is
unique and P. Helvetica occupies an isolated position in the genus (Figs P3, P4).
Description male
Measurements (n=l): Body length: 6.8 mm; wing length: 6.0 mm.
Head : Ground colour yellow. Ocellar triangle blackish. Antenna yellow, but
postpedicel mostly black, 1.2 times as long as wide at base. Arista brown, conspi-
cuously thickened at base, covered with short, thin rays. Eye roundish, small in profile
compared to gena and parafacialia. Gena about 1.2 times higher than height of eye.
Compound eye about 1.5 times higher than long. Width of frons about 1.1 times less
than its length from the two posterior ocelli to the lunule. Width of frons about 1.8
times less than width of head. Frons uniformly yellow. Gena about 1.3-1. 4 times less
high than eye. Palpus yellow, black apically. Chaetotaxy (all setae black): 2 vertical
setae; 1 fronto-orbital seta (?), postocellar seta absent.
Thorax : Ground colour yellow. Mesonotum with a broad, black stripe medially
extending almost to scutellum. A pair of lateral black stripes from level of hind margin
of the postpronotal lobe to level of dorsocentral seta. These lateral stripes may fuse
partly with the unpaired medial stripe or be interrupted on level of suture.
Subscutellum black. Anepimeron with fine, soft setulae in posterior half. Chaetotaxy
(all setae black): 1 notopleural seta; 1 supra-alar seta; 1 postalar seta; 1 dorsocentral
seta; 1 (pair of) apical scutellar seta, without soft, fine setulae along the margin.
784
A. I. SHATALKIN & B. MERZ
Legs : Generally yellow. Middle tibia with two black apical spur-like setae of
unequal size.
Wing: Entirely hyaline, with a slight yellowish tinge. Veins yellow. Section of
Ml +2 between R-M and DM-Cu about 1 .5 times as long as previous section and sub-
equal with apical section (between DM-Cu and margin). Haltere yellow.
Abdomen : Basal tergites black, other parts yellow. Terminalia not dissected.
Female
Measurements (n=2): Body length: 6. 5-7.0 mm; wing length: 7.0 mm.
Comments: Otherwise as male. Terminalia not studied.
Psila (s. str.) merdaria Collin, 1944
Material Switzerland: 1 9 , GR, 1550 m, 3 km N Lenz, 11 VII 1996, leg. B. Merz. -
1 d , TG, 460 m. Ermatingen, Neuguet, 1 V 2002, leg. K. Grimm. -19, ZH, 440 m, Ziirich-
Katzensee, 25 V 1996. leg. B. Merz.
Comments: This species is morphologically similar to P. fimetaria and it is
often difficult to separate them. It is a rather rare species which was described from
Great Britain. Since then, it has been recorded from various Central European
countries to the Moscow Region in the East and to Spain in the South (Pape, 2007;
unpublished observations).
Genus Psilosoma Zetterstedt, 1860
Type species: Scatophaga audouini Zetterstedt, 1835, by original designation.
Diagnosis: Two vertical setae. Postocellar seta absent. Notopleural seta absent.
Dorsocentral seta present or absent. 1 (pair of) apical scutellar seta. Anepimeron bare.
Hind femur in male (and also in the female of P. audouini) thickened. Male terminalia
with long aedeagus. Female ovipositor compressed laterally and strongly sclerotized.
Distribution: Two Western Palaearctic species are placed here which are
distributed in cooler regions from Northern Scandinavia southwards to the Alps where
they are usually found in higher altitudes. Both species are not uncommon in
Switzerland in higher altitudes of the Alps where they may be often swept and
observed on large stands of Alnus viridis.
Psilosoma audouini (Zetterstedt, 1835)
Material Switzerland: Id. 19, VS, 1600-2200 m, Visperterminen. Giw-Gebidem,
4 VII 2003, leg. B. Merz, J. Smit & M. van Det. - 1 9 , VS, 1900-2200 m, Visperterminen, Giw-
Gebidempass. 21 VII 2004. leg. B. Merz.
Psilosoma lefebvrei (Zetterstedt, 1835)
Material Switzerland: 1 d , GR, 2400 m, Lenzerheide, Piz Danis, 8 VIII 1992, leg. B.
Merz. - 1 d . GR. 1500 m, Valbella-Casoja, 20 VII 1996, leg. B. Merz. - 1 d . GR, 2200 m. Ftan,
Cliinas. 5 VIII 1996, leg. Merz & Bachli. - 1 9 , GR. 1300 m, Ausserferrera, 13 VIII 1991. leg.
Merz & Freidberg. - 1 9 , SZ, 1550 m. Pragelpass, 7 VIII 1995. leg. B. Merz.
Genus Chamaepsila Hendel, 1917
Type species: Musca rosae Fabricius, 1794, by original designation.
PS I U DAE OF SWITZERLAND
785
Diagnosis: Arista yellow or slightly brownish, white only in Ch. setalba
Freidberg & Shatalkin (other species with a white arista are excluded from this genus:
Psila fallax , Xenopsila spp.); postocellar seta present (usually absent in Ch. setalba):
2-3 vertical setae; 1-6 dorsocentral setae; 1 notopleural seta; 1 (pair of) apical scutellar
seta (except for Ch. obscuritarsis with 2 pairs); anepirneron entirely bare (except for
Ch.fenestralis from the Russian Far East).
Comments: Steyskal (1987) did not recognize this genus in his key to genera
and subgenera of Nearctic Psilidae. One reason may be that Psila s. str. was not yet
known from that region and he considered the subgeneric division of Hennig (1941) as
less convincing. However, taking into account the observations during the present
study and the results of the work of Buck & Marshall (2006a) this division is based on
unequivocal characters which allow its safe identification (see key above).
Chamaepsila atra (Meigen, 1826)
Material Switzerland: 2d , BE, 1430 m, Lenk. Simmenfalle, 31 V 2003. leg. B. Merz
& M. Eggenberger. - 1 9 , BE, 995 m, Tramelan, La Tourbiere. 4 VI 2003, leg. B. Merz. J.-P.
Haenni & M. Rapp. -Id, GR, 2200-2550 m, Juf-Stallerberg. 19 VI 1994, leg. B. Merz. -Id.
GR, 2250-2490 m, Lenzerheide, Piz Danis. 12 VII 1996, leg. B. Merz. - Id, GR, 1500 m.
Valbella, Casoja, 19 VII 1997, leg. B. Merz. - 1$, TI. 1965 m, Piora. 11 VI 1996, leg. M.
Bemasconi. - 1 d , 2 $ , VS, 1700-1900 m, Morgins, La Chaux-Culet, 21 VI 2003. leg. B. Merz.
- 1 d , 1 $ . VS. 1700-1950 m, Morgins, Portes du Soleil (Monthey). 22 VI 2003, leg.^B. Merz. -
1 d , VS, 1400 m, Visperterminen, Kreuz, 3 VI 2003, leg. B. Merz.
Comments: This Transpalaearctic species is characterized by the presence of 4
dorsocentral setae, this character only shared with Ch. clunalis (Collin, 1944) in the
Western Palaearctic Region. Ch. clunalis w r as described from Great Britain and was
later found in Central Europe and Spain but is still unknown from Switzerland (Pape,
2007). It differs from Ch. atra by its larger epandrium in the male which gives the
impression that the abdomen is parallel-sided (in Ch. atra more cone-like). Moreover,
the posteroventral margin of the epandrium is extended into a tooth-like projection
(Shatalkin, 1986, plate 4, fig. 9) (evenly rounded in Ch. atra). Finally, the female of C.
clunalis has therefore a wider 7th tergite which is as wide as the previous one
(decreasing in width in C. atra). C. clunalis exhibits a wade variation in colouration,
varying from black to brown, especially on the head.
Chamaepsila bicolor { Meigen, 1826)
Material Switzerland: 1 d , GR, 2100 m, Ftan-Clunas. 5 VIII 1996, leg. B. Merz & G.
Bachli. - 1 d , GR, 1660 m. S Chanf-Flin, 7 VIII 1996. leg. B . Merz & G. Bachli. - 1 $ , VS. 1550
m, Jeitzinen, 26 VI 1999. leg. B. Merz & M. Eggenberger. - 2d, VS, 1700-1900 m, Morgins,
La Chaux-Culet, 21 VI 2003, leg. B. Merz. - 2d , VS, 2040 m, Morgins, Pointe de Bellevue, 21
VI 2003. leg. B. Merz. - 1 $ , VS. 1390 m. Visperterminen/Kreuz. 3 VII 2003, leg. B. Merz. J.
Smit & M. van Det.
Comments: The specimens from "Pointe de Bellevue" are characterized by the
brown stripe on the pleura and fit therefore the description of Ch. nigromaculata
(Strobl, 1909). The two species were later synonymized (Carles-Tolra, 1993a). The
males from Switzerland correspond well w ith the illustration of the lectotype of Ch. bi-
color with the medially directed teeth on the epandrium (Soos, 1985, fig. 1 A). On the
other hand, the structure of the paramere without the tooth on the protuberance (see
786
A. I. SHATALKIN & B. MERZ
Carles-Tolra, 1993a, fig. 6) is more similar to C. quadrilineata (Strobl, 1898). Further
studies are necessary in order to establish the identity of these Swiss specimens.
Chamaepsila buccata (Fallen, 1826)
Psila gracilis Meigen, 1826; synonymy by Soos (1985).
Material Switzerland: 1 9 , GR, 680 m, Rothenbrunnen, 8 VIII 1996, leg. B. Merz &
G. Bachli. - 1 6 , 1 9 , VS, 750 m, Branson/Follateres, 29 III 2002, leg. B. Merz.
Comments: Chamaepsila buccata , Ch. sardoa (Rondani, 1876) and Ch.
rozkosnyi Carles-Tolra, 1993, form a morphologically uniform group. Ch. sardoa has
an entirely black antenna (basal segments yellowish in Ch. buccata ), 2 dorsocentral
setae (1 dorsocentral seta in Ch. buccata ), is on average smaller (3-4 mm, Ch. buccata
4-6 mm) and differs in the structure of the male terminalia (Shatalkin, 1986, plate 2,
fig. 5 for Ch. buccata (as Ch. gracilis Meigen) and plate 3, fig. 1 for Ch. sardoa). Ch.
rozkosnyi is characterized by the presence of 2 fronto-orbital setae (1 fronto-orbital seta
in Ch. buccata and Ch. sardoa) and in details of the structure of the male terminalia
with a longer aedeagus (Carles-Tolra, 1993b).
Chamaepsila buccata has a transpalaearctic distribution. Outside Europe it is
reported from Mongolia (Soos, 1974), China (Wang & Yang, 1996), and the Russian
Far East (Shatalkin, 1986).
Chamaepsila confusa sp. n. Fig. 1
Holotype: 6 , CH (=Switzerland), VS, 600 m, Leuk-Pfynwald, 25 VIII 2001 , leg. Merz
& Landry (MHNG).
Paratypes: 1 6 , same data as holotype (MHNG). -19, Switzerland, TI, 460 m, Clivio,
Arzo, 4 X 2003, LF [= Lichtfalle, = light trap], leg. H. Vicentini (MHNG). -19, France, Haute-
Savoie (74), Pormenaz, 1700-2200 m, 8-31 VII 2003, MTA [Malaise-trap], leg. Castella &
Speight (MHNG).
Diagnosis: Externally, this species is similar to Ch. persimilis (Wakerley) and
Ch. nigricornis (Meigen). It shares with Ch. persimilis the presence of a yellow spot
dorsally the anterior spiracle, but differs in the apically black palpus and the entirely
black postpedicel. Ch. nigricornis has an entirely black pleuron including the pro-
pleuron. The three species differ further in details of the male terminalia, in particular
in the shape of the aedeagus and the epandrium.
Description male
Measurements (n=2): Body length: 3.7 mm; wing length: 3.9 mm.
Head : General colour yellow but with black ocellar triangle and a pair of
narrow, black stripes on the occiput from vertex (between medial vertical setae) to the
occipital foramen. Scape and pedicel yellow, postpedicel black, about 1 .3 times as long
as basal width. Arista brownish, short setulose, longest rays shorter than diameter of
arista at base. Frons about as long as wide. Gena (in paratype) about 1 .2 times less high
than height of compound eye. Compound eye about 1.4 times as high as wide. Palpus
black apically. Chaetotaxy (all setae yellow): 3 vertical setae; 1 stout postocellar seta;
2 fronto-orbital setae.
Thorax : Almost entirely black, but with a yellow spot on propleuron dorsally of
anterior spiracle. Chaetotaxy (all setae yellow): 1 notopleural seta; 1 supra-alar seta; 1
PS I LI DAE OF SWITZERLAND
787
postalar seta; 2 dorsocentral setae posterior suture; I (pair of) apical seutellar seta.
Anepimeron bare.
Legs : Entirely yellow.
Wing\ Hyaline with yellowish veins. Section of Ml +2 between crossveins R-M
and DM-Cu about three times as long as section between BM-Cu and R-M and sub-
equal to section between DM-Cu and wing margin. Haltere yellowish white.
Abdomen: Entirely black. Male terminalia as in fig. 1 .
Female
Measurements (n=2): Body length: 4.2 mm; wing length: 4.3 mm.
Comments : Otherwise as male.
Comments: The male terminalia are characteristic for Ch. confusa. The base of
the aedeagus is convex (fig. 1), but concave in Ch. nigricornis (Lyneborg, 1964;
Shtakel'berg, 1989; van der Goot & van Veen, 1996). Ch. persimilis is characterized by
a special type of the aedeagus which has a strong desclerotization at its base in the
middle (figs 4-5).
Chamaepsila humeralis (Zetterstedt, 1847)
Material Switzerland: 1 9 , BE, 650-950 m, Mt. Raimeux, Corcelles, 2 VI 2003, leg.
B. Merz. -Id, GL, 1 130 m, Richisau (Klontal), 10 IX 1997, leg. B. Merz. - 1 6 , 1 9 , GL, 850
m, Vorauen (Klontal), 10 IX 1997, leg. B. Merz. -66 ,19 . GR, 1300-1670 m, Ausserferrera-
Cresta, 28 VIII- 1 IX 2006, leg. B. Merz. - 1 9 , GR, 980 m, Andeer (Hinterrhein), 29 VIII 2006,
leg. B. Merz. - 26 , GR, 1500 m, Valbella, Casoja, 21 VII 1997, leg. B. Merz. - 1 9 . TI, 460 m,
Clivio, Arzo, 4 X 2003, LF (= light trap), leg. H. Vicentini. - Id, VD, 1300 m, St.Cergue,
Barillette, 3 VIII 2003, B. Merz. - 2d , 1 9 , VS, 1300 m, Morgins, En Tey, 22 VI 2003, leg. B.
Merz. - 1 9 , VS, 1600-2200 m, Visperterminen,Giw-Gebidem,4 VII 2003, leg. B. Merz, J. Smit
& M. van Det.
Comments: This species is well characterized by the presence of an entirely
yellow postpronotal lobe and propleuron and thus differs from other species with
mainly black thorax and the same chaetotaxy (Ch. confusa , Ch. nigricornis , Ch. persi-
milis, Ch. rosae). The Swiss specimens show a remarkable variability in the colour of
the anepimeron. Whereas the anepimeron is usually black in this species, it is yellow
in the specimens from Morgins. The male terminalia, however, are the same in both
colour morphs and they are considered here to belong to the same species.
One male deserves a comment. It carries the following labels: "Helv., VD, 1300
m, St. Cergue, Barillette, 3 VIII 2003, leg. B. Merz". It differs from typical specimens
of Ch. humeralis by a black mesonotum with a yellow longitudinal stripe between the
postpronotal lobe and the wing base, and on the dorsal half of the pleuron. The
anepi sternum is black in the posterior half and the anepimeron slightly brownish. The
study of the male terminalia, however, does not show any difference and it is concluded
that this specimen is an extreme colour form of Ch. humeralis. A more detailed inter-
pretation of the status of this colour morph can only be obtained by the study of
additional specimens.
Chamaepsila limbatella (Zetterstedt, 1847)
Material Switzerland: 1 9 ,GR, 2100 m, Ftan-ClUnas, 5 VIII 1996, leg. B. Merz & G.
Bachli. - 1 9 , GR, 1500 m, Valbella, Casoja, 19 VII 1997, leg. B. Merz. - 1 9 , VS, 1900-
2100 m, Visperterminen, Rothorn-Giw, 30 VII 1998, leg. B. Merz & G. Bachli.
788
A. I. SHATALKIN & B. MERZ
Male terminalia of Chamaepsila spp. (1) Ch. confiisa sp. n., ventral view (holotype.CH, VS, 600
m, Leuk-Pfynwald, 25 VIII 2001). (2) Ch. pallida (Fallen), ventral view (CH, BE, Mt. Raimeux,
Corcelles. 2 VI 2003). (3) Ch. pallida (Fallen), ventral view (Russia. Moscow Region,
Golitsyno, 23 VI 1981). (4) Ch. persimilis (Wakerley), ventral view (CH, VS, Eggerberg-Erb, 24
VI 2000). (5) Ch. persimilis (Wakerley), aedeagus (F, Haute Savoie, Monnetier, Petit Saleve, 24
VIII 2003). (6) Ch. unilineata (Zetterstedt), ventral view (CH, TI, 1097 m, Mte. S. Giorgio, top,
18 V 2006). (7) Ch. unilineata (Zetterstedt). paramere, lateral view (same data).
Abbreviations: aa = aedeagal apodeme; ae = aedeagus; ep = epandrium; h = hypandrium; p =
paramere; pa = parameral arm.
PS1L1DAE OF SWITZERLAND
789
Comments: This is a rare European species which is easy to identify with its
black body and its chaetotaxy with 3 vertical setae, 1 fronto-orbital seta, and 1 dorso-
central seta (all pale).
Chamciepsila mono (Zettestedt, 1835)
Material Switzerland: 4 <3, 1 $ , BE, 1430 m, Lenk, Simmenfalle, 31 V 2003, leg. B.
Merz & M. Eggenberger. -Id, GR, 2250-2490 m, Lenzerheide, Piz Danis, 12 VII 1996. leg. B.
Merz. - 4$ , GR, 1520 m, Lenzerheide/Sundroina, 17 V 1997, leg. B. Merz. -19, GR, 1490 m,
Valbella, Casoja. 11 VII 1998, leg. B. Merz. - 2$ , VS, 1700-1900 m, Morgins, La Chaux-Culet,
21 VI 2003, leg. B. Merz. - 1 <3 , 1 9 , VS, 1700-1950 m, Morgins, Portes du Soleil (Monthey),
22 VI 2003, leg. B. Merz. - 1 9 , VS, 1900-2200 m, Visperterminen, Giw-Gebidempass, 21 VII
2004, leg. B. Merz. -39 , VS, 1600-2200 m, Visperterminen. Giw-Gebidem, 4 VII 2003, leg. B.
Merz, J. Smit & M. van Det. - 1 9 , VS, 1400 m, Visperterminen/Kreuz, 3 VI 2003, leg. B. Merz.
- 1 9 , VS, 1300-1900 m, Visperterminen, Richtung Giw, 15 VII 1995, leg. B. Merz.
Comments: Chamciepsila mono is widespread in Scandinavia and Nothem
Russia. In the South it is mainly known from higher altitudes of the Alps. In the Eastern
parts of the Palaearctic Region it is recorded from the Altai Mountains, Yakutia and
Magadan (but only females are known from the latter two localities). Soos (1974) iden-
tified this species from Mongolia based on three females.
Chamaepsila nigra (Fallen, 1820)
Material Switzerland: 1 9 , GE. 350 m, Cartigny, Moulin de Vert, 22 IV 2000, leg. B.
Merz. -19, GR, 1550 m, Lenzerheide/Sundroina, 20 VII 1997, leg. B. Merz. -19, GR, 1500
m, Valbella, Casoja, 19 VII 1997, leg. B. Merz. -19, VS, 1600-2200 m, Visperterminen, Giw-
Gebidem, 4 VII 2003, leg. B. Merz, J. Smit & M. van Det. - Id, 19, VS, 1400 m,
Visperterminen/Kreuz, 3 VI 2003, leg. B. Merz.
Chamaepsila nigricornis (Meigen, 1826)
Material Switzerland: 1 9 , BE, 650-950 m, Mt. Raimeux, Corcelles, 2 VI 2003, leg.
B. Merz. - 2d, GE, 420 m, Bernex, Bois de Carabot, 29 IV 2005, leg. B. Merz. - 29 , GE,
510 m, Bernex-Signal, 24 IV 2003, leg. B. Merz & F. Amiet. -Id, GE, 500 m. Bernex-Signal,
30 IV 2007, leg. B. Merz. -19, GE, Avusy, Moulin de la Grave, 15 V 2003, leg. H. Boillat. -
Id, GR, 1600 m. Lenzerheide/Sundroina. 16 VII 2000. leg. B. Merz. - Id, 29, GR. 1300-
1325 m, Ausserferrera, 28-30 VIII 2006, leg. B. Merz. - 1 9 , JU, 1000 m, Le Chaux, Les
Breuleux, La Tourbiere, 28 VI 2003, leg. B. Merz. - 1 9 , TI, 370m, Cugnasco-Montedato, 2 IV
2007. B. Merz. - 1 9 ,TI, 205 m, Gordola, Verzascaufer, 1 IV 2007, leg. B. Merz. -19, VS, 650
m, Baltschieder, Chumme, 23 V 1998, leg. B. Merz. - Id, VS, 920 m, Leuk-Brentjong, 15 V
2000, leg. B. Merz. - Id, VS, 625 m, Leuk-Platten, 2 V 1999, leg. B. Merz. - 1 9 , VS. 1700-
1900 m, Morgins, La Chaux-Culet, 21 VI 2003, leg. B. Merz. - 2d, VS, 1550 m,
Visperterminen, ob Dorf, 27 VIII 2001 , leg. B. Merz & B. Landry. - 29 , ZH. 430 m, Ziirich-
Allmend, 6 V 1998, leg. B. Merz. - 1 9 ,ZH, 630 m, Zurich-Zurichberg, 6 IX 1992, leg. B. Merz.
Comments: Chamaepsila nigricornis is known as a pest of Chrysanthemum sp.
with its larvae feeding in the roots (Vernon, 1962; Iwasa, 1998). Morphologically, it
can be confused with Ch. confusa , Ch. persimilis , and Ch. rosae which have the same
general body colouration and chaetotaxy. However, their male terminalia are different
and allow an easy identification (Wakerley, 1959; Lyneborg, 1964; Wang, 1988;
Shtakel'berg, 1989).
Chamaepsila obscuritarsis (Loew, 1856)
Material Switzerland: Id, ZH. 650 m, Zurich-Zurichberg. 16-20 VI 1995, MF
[=Malaisefalle], leg. S. Ungricht.
790
A. I. SHATALKIN & B. MERZ
Comments: With its two (pairs of) scutellar setae this species is unique within
Chamaepsila. For this reason Frey (1925) proposed the genus Tetrapsila for it with the
remark that this genus may be only a subgenus of Chamaepsila . The two genera were
indeed synonymized later (Shatalkin, 1986; Iwasa, 1998).
Chamaepsila pallida (Fallen, 1820) Figs 2-3, P5-P6
Material Switzerland: 1 6 , 1 9 , BE, 650-950 m, Mt. Raimeux. Corcelles, 2 VI 2003,
leg. B. Merz. - 1 9, BE, Burgdorf, leg. Meyer-Dtir (small specimen). - 2 9 , GR, 1530-1540 m,
Valbella, Casoja, 14-15 VII 1998, leg. B. Merz. - 1<3, GR, 1530 m, Valbella. Casoja. 19 VII
1997, leg. B. Merz. -19, GR, 1430 m, Zemez, Gondas, 6 VIII 1996, leg. B. Merz & G. Bachli.
-39, VS, 1400 m. Visperterminen, 14 VII 1995, leg. B. Merz.
Diagnosis: Ch. pallida is morphologically similar with Ch. luteola (Collin,
1944). They have both a yellow body. The latter differs from Ch. pallida by other pro-
portions of its eye, the brownish dorsal border of the postpedicel (uniformly yellow in
Ch. pallida), the brownish yellow setae on head and thorax (yellow in Ch. pallida) and
the structure of the male terminalia (Wang, 1988). Two other species, Ch. nigrosetosa
Frey, 1925, and Ch. unilineata (Zetterstedt, 1847), were often synonymized with Ch.
pallida. According to our studies we treat them here as good species. They may be
separated from each other with the key provided at the end of this chapter.
Comments: K. Holston from the NRS kindly made available photos of the
entire type series of Scatophaga pallida Fallen (figs P5-P6). It contains one male, six
females, and one specimen without abdomen. The male (fig. P5) and one female (fig.
P6) carry an identification label in Fallen's handwriting. The male is labelled "S.
pallida" (underlined), 6 (written on the second line). The type locality is according to
the original description "Esperod" in Southern Sweden (Fallen, 1820). The specimen
without abdomen belongs to another species, probably Chamaepsila rufa (Meigen,
1826), with its black setae on head and thorax. The abdomen is darkened in two
females, and one of them has also the tip of the palpus darkened. They correspond
probably to the description of Var. (3 of Fallen. The male and the other four females are
yellow including the palpus and are considered here as syntypes of Ch. pallida Fallen
s. str. A short description of the male syntype (fig. P5) based partly on the study of the
photos is provided here.
Description male
Head: Mostly yellow, but ocellar triangle blackish. Antenna entirely yellow,
postpedicel about 1.6 times as long as wide at base. Arista yellowish, short setulose.
Gena about 1.2 times less high than eye. Palpus yellow. Chaetotaxy (all setae light
yellowish): 3 vertical setae; 1 stout postocellar seta; 2 fronto-orbital setae.
Thorax : Yellow including subscutellum. Chaetotaxy (all setae light yellowish):
1 notopleural seta; 2 dorsocentral setae posterior of suture; 1 (pair of) apical scutellar
seta.
Legs: Yellow.
Wing: Flyaline with yellowish veins. Section of Ml +2 between R-M and DM-
Cu about 2.9 times as long as previous section (between BM-Cu and R-M) and slightly
longer than last section (between DM-Cu and wing margin). Haltere yellowish white.
Abdomen: Entirely yellow. Genitalia (figs 2-3) with a wide aedeagus; paramere
without a distinct, large tooth.
PS I LI DAE OF SWITZERLAND
791
Chamaepsila pectoralis (Meigen, 1826)
Material Switzerland: 1 9 , GR, 980 m, Andeer-Clugin. 17 VI 1994, leg. B. Merz. -
1 9 , GR, 1490 m, Valbella-Casoja, 26 Vll 1999, ZF, leg. B. Merz. - 1 d , SG, 610 m, Wattwil, 1 1
VII 1997, leg. B. Merz. - 1 9 , ZH, 400 m, Embrach-Haumuli, 4 VI 1998, leg. C. Wolf.
Comments: The specimen from Valbella was collected in a tent-trap.
Chamaepsila persimilis (Wakerley, 1959) Figs 4-5
Material Switzerland: 1 d, GE, Corsier-Port, vitre veranda, 3-15 VIII 2003, leg. C.
Besuchet. - 1 <3, 19, GE, 420 m, Dardagny/Roulave, 30 VI 2001, leg. B. Merz. - Id, GE,
360 m, Dardagny, Le Moulin, 30 VI 2001 . leg. B. Merz. -19, GE, 350 m, Cartigny, Moulin de
Vert, 5 VI 2006, leg. B. Merz. -Id, GE, 360 m, Dardagny, Le Moulin, 8 VII 2002, leg. B. Merz.
-19.GR, 980 m. Andeer (Hinterrhein), 29 VIII 2006, leg. B. Merz. - 5 9 , GR, 350 m, Grono
(Dorf), 31 VIII 2006, leg. B. Merz. - 29, GR, 330 m, Grono (Moesa), 31 VIII 2006, leg. B.
Merz. - 19, SG, 610 m, Wattwil, 11 VII 1997, leg. B. Merz. - 2d, 19, VS, 460-750 m,
Branson/Follateres, 9 VI 2004, leg. B. Merz & J.-P. Haenni. -19, VS, 1550 m, Visperterminen,
ob Dorf, 27 VIII 2001 , leg. B. Merz & B. Landry. - 2d , VS, 920 m, Leuk-Brentjong, 15 V 2000,
leg. B. Merz. - Id, 29, VS, 745 m, Visp, Hohbrunne, 3 VI 2003, leg. B. Merz. - Id, ZH,
530 m, Ziirich-Honggerberg, 16 VI 1998, leg. B. Merz.
Comments: The aedeagus of this species is unique. According to the available
illustrations (Wakerley, 1959; Lyneborg. 1964; van der Goot & van Veen, 1996) and
our observations (figs 4-5) it has a deep incision at its base which may be interpreted
as a desclerotized area. In contrast, the lateral basal protuberances are strongly sclero-
tized. Moreover, the epandrium is unique by its extended and apically narrowed pos-
teroventral margin (fig. 4).
Chamaepsila quadrilineata (Strobl, 1898)
Material Switzerland: 1 9, VS, Jeitzinen, Agerde, 3 VII 2001, leg. B. Merz. - 3d,
2 9 , VS. 1700-1900 m, Morgins, La Chaux-Culet, 21 VI 2003, leg. B. Merz. - 3 9 , VS, 2040 m,
Morgins, Pointe de Bellevue, 21 VI 2003, leg. B. Merz.
Chamaepsila rosae (Fabricius, 1794)
Material Switzerland: ld,GR, 1520 m, Lenzerheide-Sundroina, 12 VII 1999, leg. B.
Merz. - Id, GR, 1490 m, Valbella-Casoja, 24 VII 1999, leg. B. Merz. - Id, VS, 700-770 m,
Raron, Heidnischbiel, 26 V 1999, leg. B. Merz. - Id, VS, 1600-2200 m, Visperterminen, Giw-
Gebidem, 4 VII 2003, leg. B. Merz.
Comments: This is the noxious "carrot fly" whose larvae are tunneling roots of
carrot, celery, parsnip, parsley and other umbelliferous plant species (Grichanov et al.,
2005), It is widely distributed in the Palaearctic Region. In the Eastern Palaearctic
Region it is recorded from China (Wang & Yang, 1996), Mongolia (Soos, 1974), the
Amur area (Shatalkin, 1986), and the Sakhalin and Kuril Islands. Surprisingly it has
not yet been found in the Primorye Territory of Russia, maybe because this species is
replaced there by the morphologically (and maybe biologically) similar species Ch.
sonora (Shatalkin, 1986) and Ch. iwasai (Shatalkin, 1996) which were found in a
garden of a kitchen. Although Ch. rosae is known from Japan it is rare because another
species, Synaphopsila nartshukae Shatalkin, 1986, is known to damage carrots (Iwasa
et al ., 1987; Narita et al ., 2005). Later, the species was accidently introduced to North
America where it is now a pest species (Hooper et al ., 2002) and to New Zealand
(Evenhuis, 1989).
792
A. I. SHATALKIN & B. MERZ
The name Musca rosae Fabricius, 1794, is a primary homonym of Musca rosae
De Geer, 1776 (a junior synonym of the Syrphidae Musca pyrastri Linnaeus, 1758)
(Thompson & Pont, 1994) and the new name Chamaepsila hennigi Thompson & Pont,
1994, was proposed for the carrot fly. Later, Chandler (1998b) requested the ICZN to
preserve the name "rosae" for the carrot fly, and this proposition was accepted (ICZN,
1999).
The identity of the Swiss specimens was confirmed by dissection of the male
terminalia and the comparison with illustrations published by Wakerley (1959),
Lyneborg (1964) and Wang (1988).
Chamaepsila sardoa (Rondani, 1876) - new for Switzerland
Material Switzerland: 19. TI, 205 m, Gordola, Verzascaufer, 15 V 2006, leg. B.
Merz. - 1 d . 1 9 . TI. 205 m. Gordola, Verzascaufer, 1 IV 2007. leg. B. Merz.
Comments: This species is in most respects similar to Ch. buccata. It may be
separated from the latter and from Ch. rozkosnyi , another morphologically similar
species, by the characters mentioned under Ch. buccata. Ch. sardoa is a Mediterranean
species which has been found from Spain to Israel. The present finding in Southern
Switzerland represents the northernmost record of this species. Outside the Western
Palaearctic Region it was recorded from desert zones in Middle Asia and from
Mongolia (Soos, 1974). The record from Myanmar by Frey (1955) is based on a
misidentification. He compared his female from Mt. Kambaiti with a female from
Morocco which he thought was Ch. sardoa. A careful study of both specimens revealed
that they belong to two different species. The female from Morocco was later described
as Psila atlasica Shatalkin, 2000. and the female from Myanmar as Psila freyi
Shatalkin, 2000.
Chamaepsila unilineata (Zetterstedt, 1847) stat. rev. - new for Switzerland Figs 6-7
Material Switzerland:
A) Typical specimens: Id, GR, 1000-1150 m, Brienz-Surava, 25 VII 1999, leg. B.
Merz. -19, GR. 1550 m, Lenzerheide/Sundroina, 20 VII 1997, leg. B. Merz. - 6d, 39 , GR,
1520 m, Lenzerheide/Sundroina, 10 VII 1998. leg. B. Merz. - 6 d, 29, GR, 1490 m,
Lenzerheide/See, 1 1 VII 1998. leg. B. Merz. -19. GR. 1500 m. Valbella. Casoja, 21 VII 1997,
leg. B. Merz. -19, GR. 1530 m, Valbella. Casoja. 14 VII 1998, leg. B. Merz. - 2d , 39 , GR,
1490 m. Valbella, Casoja, 24 VII 1999. leg. B. Merz. -49 , TI, 1097 m. Mte. S. Giorgio, top, 18
V 2006, leg. B. Merz. - 6d, 39, VS, 1550 m, Jeitzinen, 26 VI 1999, leg. B. Merz & M.
Eggenberger. - 4d , 3 9 , VS. 1500 m, Jeitzinen. Agerde, 3 VII 2001 . leg. B. Merz. - 1 d . VS.
1400 m. Visperterminen. 14 VII 1995, leg. B. Merz.
B) Longitudinal stripes on mesonotum badly expressed or absent: 1 d . GR. 980 m.
Andeer, Clugin, 17 VI 1994, B. Merz. - 29, GR. 1550 m, 3 km N Lenz, 11 VII 1996, leg. B.
Merz. - 2d, 29. GR, 1520 m, Lenzerheide/Sundroina, 10 VII 1998. leg. B. Merz. - 19, GR.
1600 m. Lenzerheide/Sundroina. 16 VII 2000. leg. B. Merz. - 5d, 19, GR, 1490 m,
Lenzerheide/See, 1 1 VII 1998, leg. B. Merz. -Id, GR. 1500 m, Valbella. Casoja, 19 VII 1997.
leg. B. Merz. - 3d . 29 . GR, 1490 m, Valbella. Casoja. 24 VII 1999, leg. B. Merz. - 2d, 1 9 ,
GR. 1000-1150 m, Brienz-Surava, 25 VII 1999, leg. B. Merz.- 2d, TI. 1097 m.Mte. S. Giorgio,
top. 18 V 2006. B. Merz. -19, ZH. 350 m. Flaach. Thurauen. 6 VI 1993, leg. B. Merz.
C) Katepisternum brownish: 1 9 , GR, 1520 m, Lenzerheide/Sundroina. 10 VII 1998.
leg. B. Merz. - Id. GR. 1490 m. Valbella. Casoja. 24 VII 1999, leg. B. Merz. - 5d, 49 , VS,
1550 m, Jeitzinen, 26 VI 1999, leg. B. Merz & M. Eggenberger. - 2d. 19, VS. 1500 m.
Jeitzinen. Agerde. 3 VII 2001, leg. B. Merz.
PS I LI DAE OF SWITZERLAND
793
Comments: So far, this species has usually been synonymized with
Chamaepsila pallida (Fallen). It was argued that the presence of three dark longitu-
dinal stripes on the yellow mesonotum and the dark subscutellum fall within the colour
variation of Ch. pallida (Hennig, 1941; Soos, 1984, 1985). However, a careful study
including the structure of the male terminalia has shown that two species are involved.
They differ in the colour of the subscutellum which is yellow in Ch. pallida but has a
brown median stripe in Ch. unilineata A brief diagnosis of Ch. unilineata is provided
here.
Description Male
Head: Antenna with postpedicel yellow, rarely with a narrow or wide brownish
dorsal margin. Arista rather long setulose (as in Hennig, 1941, fig. 19, right illus-
tration); Palpus yellow.
Thorax : Mesonotum yellow, with a medial and a pair of lateral brown to black,
narrow to wide stripes which may be conspicuous or more or less reduced or even
almost absent. Subscutellum at least medially brown to black. Katepisternum yellow
or with brown spot of various size.
Male terminalia: Aedeagus rather narrow, triangular. Paramere with four large
teeth (figs 6-7).
Chamaepsila villosula (Meigen, 1826)
Material Switzerland: 46 . GR, 2100 m, Ftan/Clunas, 5 VIII 1996, leg. B. Merz & G.
Bachli. - 1 6 , VS, 1500 m, Jeitzinen, Agerde, 3 VII 2001, leg. B. Merz.
Comments: This species is easy to identify because of its colouration, chaeto-
taxy and the male terminalia with the large epandrium. It is rarely collected and its
distribution is therefore little known. Pape (2007) recorded it from Belgium, Germany,
Czech and Slovak Republics, Hungary, and Switzerland. Recently, it was found in Italy
(Verona province, Chiesa S. Michele, 26 VI 2001, leg. B. Merz & F. Mason, MHNG)
and in France (Hautes-Alpes, Montgenevre, 1800 m, VII. leg. M. Bartak, ZMUM).
TAXONOMIC REMARKS CONCERNING THE CHAMAEPSILA PALLIDA GROUP
Frey (1925) attempted to classify the species of Chamaepsila into more or less
homogenous, "natural" groups based on their chaetotaxy. His "Gruppe 3", here called
Chamaepsila pallida group, is characterized by the presence of 3 vertical setae, 2 dor-
socentral setae posterior the suture and 1 well-developed postocellar seta. Body colour
varies from almost entirely black {Ch. rosae subgroup, including Ch. confasa , Ch.
humeralis , Ch. nigricornis , Ch. persimilis, Ch. rosae) to partly yellow and partly black
{Ch. pectoralis subgroup with Ch. pect oralis and Ch. strigata) to almost entirely
yellow {Ch. pallida subgroup). The latter subgroup is taxonomically the most difficult
one. The following seven species which are recognized here as valid are assigned to
this subgroup: Ch. andreji (Shatalkin, 1996), Ch. luteola (Collin, 1944), Ch. nigro -
setosa Frey, 1925, Ch. ozerovi (Shatalkin, 1993), Ch. pallida (Fallen, 1820), Ch. trior -
biseta Papp, 2003, and Ch. unilineata (Zetterstedt, 1847). Two of these species, Ch. ni-
grosetosa and Ch. unilineata , are subject to various interpretations by different
students of the family and their status is revised here based on the study of additional
specimens.
794
A. I. SHATALKIN & B. MERZ
Chamaepsila nigrosetosa was described from one female from Finland. Hennig
(1941) considered it as a colour morph of Ch. pallida , and Soos (1984) followed this
interpretation. However, it differs from the latter clearly in several external characters
and this may be the reason why it was recognized as a good species in the recent
checklist of Diptera of Czech and Slovak Republics (Bartak & Carles-Tolra, 2006) and
in the Fauna Europaea database (Pape, 2007).
The status of Ch. unilineata was often questioned in the past, and it was usually
treated as a colour form of Ch. pallida (see above, under this species). Zetterstedt
(1847) provided a rather good description which allows to differentiate it from the
other species of Chamaepsila known at that time, especially from Ch. pallida.
Unfortunately, Wang (1988) in her revision of the West Palaearctic Chamaepsila ,
dissected only one male of the "typical" Ch. pallida , but none of the form with black
stripes {"Ch. unilineata ") and she treated Ch. unilineata as a variety of Ch. pallida in
the key. On the other hand, Bartak & Carles-Tolra (2006) and Pape (2007) considered
both species as valid. It should be mentioned here that Seguy (1934) distinguished the
two species in his key, one as Ch. pallida, the other as Ch. debilis (Egger, 1862). From
his descriptions it is evident that his "Ch. debilis" corresponds with the original
description of Ch. unilineata. He thought to follow basically the interpretation of
Schiner (1863) who included the two species {pallida and debilis) in his key of
Austrian Diptera. However, the description of "Psila debilis" sensu Schiner
corresponds better with the current interpretation of Ch. nigrosetosa than of Ch. unili-
neata (which was diagnosed by Schiner in a footnote). The problems concerning the
status of Ch. unilineata may be explained by the high variability of its external
characters, in particular the development of the longitudinal, black, narrow stripes on
the mesonotum which may be conspicuous, faint or absent (see "Material" above under
Ch. unilineata). However, based on a careful study of numerous specimens it is
concluded here that both species are valid and may be separated based on a combi-
nation of external and genitalian characters. In order to facilitate identification, an
updated key for species of the Ch. pallida group is provided here.
A SHORT KEY TO SPECIES OF THE EUROPEAN SPECIES OF THE CHMAE -
PSILA PALLIDA GROUP
This key is based basically on external characters. Illustrations of the male
terminalia may be found in Shatalkin (1986) and Wang (1988) for most species
described until then and in the original descriptions for the species described since
then. The species marked with an asterisk (*) were studied by us. The other species
were included in the key based on their original descriptions.
1 a Mesonotum entirely yellow 2
lb Mesonotum yellow with black stripes or entirely black 7
2a Subscutellum yellow. Arista short setulose, rays shorter than basal
diameter of arista 3
2b Subscutellum brown or black at least medially. Arista long setulose, rays
longer than basal diameter of arista (see Hennig 1941, fig. 19, right
illustration) Chamaepsila unilineata (Zetterstedt, 1847) partim*
(= Ch. pallida auct. nee. Fallen)
PSIUDAE OF SWITZERLAND
795
3a Abdominal tergites black. Palpus apically black 4
3b Abdominal tergites yellow. Palpus yellow, or narrowly brownish apically ... 6
4a 3 fronto-orbital setae present. Frons yellow, with a transverse black
stripe dorsally the lunule. Body length: 3.45 mm. - Hungary
Chamaepsila triorbiseta Papp, 2003
4b 1-2 fronto-orbital setae present. Frons yellow throughout 5
5a Postpedicel black. Setae on head and thorax ranging from yellow (spe-
cimens from the Caucasus) to dark brown (specimens from Northern
Italy). Body length: 3.3 mm. - Russia (Caucasus), Italy
Chamaepsila andreji (Shatalkin, 1996)*
5b Postpedicel mainly yellow, with a darkened dorsal margin. Setae on head
and thorax black. Body length: 4.5 mm. - Finland, Czech and Slovak
Republics, Russia (Moscow area) Chamaepsila nigrosetosa Frey, 1925*
6a Postpedicel yellow. Setae on head and thorax light yellow. Body length:
4.0-5 .0 mm. - Europe, Mongolia (?).... Chamaepsila pallida (Fallen, 1820)*
6b Postpedicel mainly yellow, with a darkened dorsal margin. Palpus often
narrowly brownish apically. Setae on head and thorax brownish yellow.
Body length: 4.0 mm. - Great Britain, Denmark, Czech and Slovak
Republics, Russia (Moscow area) Chamaepsila luteola (Collin, 1944)*
7a Mesonotum yellow, with 3 longitudinal stripes. Pleuron yellow.
Postpedicel mainly yellow, sometimes with a darkened dorsal margin.
Arista long setulose, rays longer than basal diameter of arista (see
Hennig 1941 , fig. 19, right illustration). Male: Aedeagus narrow and pa-
ramere with four large teeth (figs 6-7). Body length: 4.0-5 .0 mm -
Europe Chamaepsila unilineata (Zetterstedt, 1847) partim*
7b Mesonotum entirely black. Other characters variable 8
8a Pleuron black, or black with a yellowish spot dorsally the anterior
spiracle 9
8b Pleuron partly or entirely yellow, at least postpronotal lobe yellow 12
9a Pleuron with a yellowish spot dorsally the anterior spiracle. Praelabrum
yellow 10
9b Pleuron entirely black. Praelabrum black 11
10a Palpus yellow, rarely brownish infuscated. Postpedicel black but yellow
ventrally at base. Male: terminalia as in figs 4-5. Body length: 3.5-
3.8 mm. - Europe Chamaepsila persimilis (Wakerley, 1959)*
10b Palpus black in apical half. Postpedicel entirely black. Male: terminalia
as in fig. 1. Body length: 3.7-4. 2 mm. - France, Switzerland
Chamaepsila confusa sp. n.*
1 la Postpedicel black, but with a small yellow spot dorsally at base around
and/or basally the insertion of the arista. Body length: 3.2-3 .8 mm. -
Holarctic species Chamaepsila rosae (Fabricius, 1794)*
lib Postpedicel entirely black Body length: 3.4-4 .2 mm. - Palaearctic,
Oriental (Myanmar), Afrotropical (Kenya?) Regions
Chamaepsila nigricornis (Meigen, 1826)*
796
A. I. SHATALKIN& B. MERZ
12a Pleuron black, but postpronotal lobe (always) and sometimes also
anepimeron yellow. Body length: 2.8-3 .7 mm. - Europe
Chamaepsila humeralis (Zetterstedt, 1847)*
12b At least ventral half of pleuron yellow 13
13a Pleuron entirely yellow. Body length: 2.7-3. 2 mm. - Europe
Chamaepsila pectoralis (Meigen, 1826)*
13b Pleuron bicoloured, dorsal half black, ventral half yellow. Body length:
3.0 mm. - Montenegro Chamaepsila strigata (Collin, 1959)
CONCLUSIONS
The present paper summarizes the present knowledge of the family Psilidae for
Switzerland and discusses some taxonomical problems related to the European fauna.
Up to 1998 reliable data for 6 species only were published from Switzerland (Ringdahl,
1957). They were all discovered after 1990 again. Whereas 32 species have been
recorded from Switzerland in the checklist and its first supplement (Merz, 1998; Merz
et al ., 2002). another 4 species are added here including two species new to science.
Currently, 36 species are known with certitude from this country. Based on the
checklists of other Central European countries (Germany, the Netherlands, Czech and
Slovak Republics, Hungary) and our knowledge of the general distribution of the
species it may be expected that another 6 species at least may be found with more
extensive field work focused on this family. They are often either rare in collections,
they have been overlooked due to their similarity with common species, or they were
misidentified by non-expert entomologists. This list of expected species comprises
Chamaepsila andreji (Shatalkin, 1996), Ch. clanalis (Collin, 1944), C/z. luteola
(Collin, 1944), Ch. rozkosnyi Carles-Tolra, 1993, Ch. rufa (Meigen, 1826), and C/z tri-
orbiseta Papp, 2003. For instance, Ch. andreji and C/z. rozkosnyi were recently disco-
vered in Northern Italy less than 50 km from the Swiss border, Ch. clunalis and C/z.
luteola , both described from Great Britain, have a large distribution in Europe, as it is
the case for C/z. rufa. On the other hand, C/z. triorbiseta was described few years ago
from a single female only. It seems to be a rare species and may require special
collecting techniques. The peculiar geographical position of Switzerland in the center
of Europe with influences from various climatic areas allowed the occurrence of a
diverse fauna with elements from different parts of Europe. The following three major
elements constitute the Psilidae fauna of Switzerland. The largest number is represen-
ted by species which have a large distribution in Europe or even the entire Palaearctic
Region (for instance, Chamaepsila nigricornis. Ch. rosae , Psila fimetaria) . The second
element comprise boreal and subarctic species of higher altitudes (generally the Alps)
in Switzerland, like the two species of Psilosoma, Chamaepsila atra , or C/z. morio.
Finally, Chamaepsila sardoa is recorded here for the first time from Southern
Switzerland which represents the northernmost record of this Mediterranean element.
A new insight into some difficult species groups of Chamaepsila is presented
here. In particular, the status of C/z. unilineata is revised based on numerous specimens
from Switzerland. They allow to conclude that it is a good species clearly different
from C/z. pallida with which it was usually synonymized. An updated key should
permit to identify this and the other species of the C/z. pallida group with more confi-
PS 1 LI DAE OF SWITZERLAND
797
dence. However, additional work is needed in order to understand fully the taxonomy
of the difficult genus Chamaepsila. In particular, the study of some types should allow
to fix the status of some of the enigmatic species which are known from few specimens
or the type series only.
ACKNOWLEDGEMENTS
We wish to thank the following colleagues for much helpful advise and assis-
tance during the preparation of this study (in alphabetical order): Dr. M. Bartak (Czech
University of Agriculture, Prague, Czech Republic), Dr. A. Freidberg (Tel-Aviv
University, Tel-Aviv, Israel), Dr. Kevin Holston (Swedish Museum of Natural History,
Stockholm, Sweden), Dr. Vladislav Martinekt, Dr. A. L. Ozerov (Zoological Museum,
Moscow State University, Russia) and Dr. Laszlo Papp (Hungarian Natural History
Museum. Budapest, Hungary). The second author is indebted to many colleagues for
their company during the numerous field trips and for the specimens they put at our
disposal (see under "Material " for their names). For technical help we express our best
thanks to Florence Marteau, Corinne Reuteler, and Philippe Wagneur (MHNG).
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APPENDIX
A REVISED CHECKLIST OF THE PSILIDAE OF SWITZERLAND
The following abbreviations are used behind the names: N = New record for Switzerland in the
present paper; R1 = Merz (1998); R2 = Merz et al. (2002)
Subfamily Chylizinae
Chyliza Fallen, 1820
- annulipes Macquart, 1835 R1
- extenuata (Rossi, 1790) R1
- leptogaster (Panzer, 1798) R1
- nova Collin, 1944 R1
- vittata Meigen, 1826 R1
Subfamily Psilinae
Tribe Loxocerini
Imantimyia Frey, 1925
- albiseta (Schrank, 1803) R1
- fulviventris (Meigen, 1826) R1
- nigrifrons (Macquart, 1835) R1
- sylvatica (Meigen, 1826) R1
Loxocera Meigen, 1803
subgenus Loxocera Meigen, 1803
- aristata (Panzer, 1801) R1
- maculata Rondani, 1876 R2
subgenus Platystyla Macquart, 1835
- hojfmannseggi Meigen, 1826 R1
Tribe Psilini
Chamaepsila Hendel ,1917
- atra (Meigen, 1826) R1
- bicolor (Meigen, 1826) R1
- buccata (Fallen, 1826) R1
- confusa Shatalkin & Merz, sp. n. N
- burner alis (Zetterstedt, 1847) R1
- limbatella (Zetterstedt, 1847) R1
- morio (Zetterstedt, 1835) R1
- nigra (Fallen, 1820) R1
- nigricornis (Meigen, 1826) R1
- obscuritarsis (Loew, 1856) R1
- pallida (Fallen, 1820) R1
- pectoralis (Meigen, 1826) R1
- persimilis (Wakerley, 1959) R1
- quadrilineata (Strobl, 1898) R1
- rosae (Fabricius, 1794) R1
- sardoa (Rondani, 1876) N
- unilineata (Zetterstedt, 1847) N
- villosula (Meigen, 1826) R1
Psila Meigen, 1803
subgenus Oxypsila Frey, 1925
- abdominalis Schummel, 1844 R1
subgenus Psila Meigen, 1803
-fimetaria (Linnaeus, 1761 ) R 1
- helvetica Shatalkin & Merz, sp. n. N
- merdaria Collin, 1944 R1
Psilosoma Zetterstedt, 1860
- audouini (Zetterstedt, 1835) R1
- lefebvrei (Zetterstedt, 1835) R1
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